Hyena

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Hyenas
Temporal range: 22–0  Ma
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Early Miocene – recent
Hyaenidae Diversity.jpg
The four living species of hyena, clockwise from upper left: spotted hyena ( Crocuta crocuta ), brown hyena ( Hyaena brunnea ), aardwolf ( Proteles cristata ) and striped hyena ( Hyaena hyaena )
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Infraorder: Viverroidea
Family: Hyaenidae
Gray, 1821
Genera
Hyaenidae range.png
Synonyms
Aardwolf, smallest member of the Hyena family, skeleton. (Museum of Osteology) Aardwolf Skeleton.jpg
Aardwolf, smallest member of the Hyena family, skeleton. (Museum of Osteology)

Hyenas or hyaenas (from Ancient Greek ὕαινα, hýaina [1] ) are feliform carnivoran mammals of the family Hyaenidae /hˈɛnɪd/ . With only four extant species (in three genera), it is the fifth-smallest biological family in the Carnivora, and one of the smallest in the class Mammalia. [2] Despite their low diversity, hyenas are unique and vital components of most African ecosystems. [3]

Contents

Although phylogenetically closer to felines and viverrids, as part of suborder Feliformia, hyenas are behaviourally and morphologically similar to canines in several elements of convergent evolution; both hyenas and canines are non-arboreal, cursorial hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable claws are adapted for running and making sharp turns. However, hyenas' grooming, scent marking, defecation habits, mating and parental behaviour are consistent with the behaviour of other feliforms. [4]

Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children. [5] Other cultures associate them with witchcraft, using their body parts in traditional African medicine. [6]

Evolution

Origins

Hyenas originated in the jungles of Miocene Eurasia 22 million years ago, when most early feliform species were still largely arboreal. The first ancestral hyenas were likely similar to the modern African civet; one of the earliest hyena species described, Plioviverrops , was a lithe, civet-like animal that inhabited Eurasia 20–22 million years ago, and is identifiable as a hyaenid by the structure of the middle ear and dentition. The lineage of Plioviverrops prospered, and gave rise to descendants with longer legs and more pointed jaws, a direction similar to that taken by canids in North America. [7] Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15 million years ago (with one taxon having colonised North America), they became extinct after a change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous aardwolf survived, while the bone-crushing hyenas (including the extant spotted, brown and striped hyenas) became the undisputed top scavengers of Eurasia and Africa. [7]

Rise and fall of the dog-like hyenas

Skull of Ictitherium viverrinum, one of the "dog-like" hyenas. American Museum of Natural History Ictitherium viverrinum.JPG
Skull of Ictitherium viverrinum, one of the "dog-like" hyenas. American Museum of Natural History

The descendants of Plioviverrops reached their peak 15 million years ago, with more than 30 species having been identified. Unlike most modern hyena species, which are specialised bone-crushers, these dog-like hyenas were nimble-bodied, wolfish animals; one species among them was Ictitherium viverrinum, which was similar to a jackal. The dog-like hyenas were very numerous; in some Miocene fossil sites, the remains of Ictitherium and other dog-like hyenas outnumber those of all other carnivores combined. The decline of the dog-like hyenas began 5–7 million years ago during a period of climate change, which was exacerbated when canids crossed the Bering land bridge to Eurasia. One species, Chasmaporthetes ossifragus , managed to cross the land bridge into North America, being the only hyena to do so. Chasmaporthetes managed to survive for some time in North America by deviating from the cursorial and bone-crushing niches monopolised by canids, and developing into a cheetah-like sprinter. Most of the dog-like hyenas had died off by 1.5 million years ago. [7]

Bone-crushing hyenas

By 10–14 million years ago, the hyena family had split into two distinct groups: dog-like hyenas and bone-crushing hyenas. The arrival of the ancestral bone-crushing hyenas coincided with the decline of the similarly built family Percrocutidae. The bone-crushing hyenas survived the changes in climate and the arrival of canids, which wiped out the dog-like hyenas, though they never crossed into North America, as their niche there had already been taken by the dog subfamily Borophaginae. By 5 million years ago, the bone-crushing hyenas had become the dominant scavengers of Eurasia, primarily feeding on large herbivore carcasses felled by sabre-toothed cats. One genus, Pachycrocuta , was a 200 kg (440 lb) mega-scavenger that could splinter the bones of elephants [ citation needed ]. With the decline of large herbivores by the late ice age, Pachycrocuta was replaced by the smaller Crocuta. [7]

Rise of modern hyenas

Skeletons of a striped hyena and a spotted hyena, two species of the "bone-crushing" hyenas Stripedspottedhyenas.jpg
Skeletons of a striped hyena and a spotted hyena, two species of the "bone-crushing" hyenas

The four extant species are the striped hyena (Hyaena hyaena), the brown hyena (Hyaena brunnea), the spotted hyena (Crocuta crocuta), and the aardwolf (Proteles cristata).

The aardwolf can trace its lineage directly back to Plioviverrops 15 million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. Its unrivaled ability to digest the terpene excretions from soldier termites is likely a modification of the strong digestive system its ancestors used to consume fetid carrion. [7]

The striped hyena may have evolved from H. namaquensis of Pliocene Africa. Striped hyena fossils are common in Africa, with records going back as far as the Villafranchian. As fossil striped hyenas are absent from the Mediterranean region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of spotted hyenas in Asia at the end of the Ice Age. The striped hyena occurred for some time in Europe during the Pleistocene, having been particularly widespread in France and Germany. It also occurred in Montmaurin, Hollabrunn in Austria, the Furninha Cave in Portugal and the Genista Caves in Gibraltar. The European form was similar in appearance to modern populations, but was larger, being comparable in size to the brown hyena. [8]

The spotted hyena (Crocuta crocuta) diverged from the striped and brown hyena 10 million years ago. [9] Its direct ancestor was the Indian Crocuta sivalensis, which lived during the Villafranchian. [10] Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp carnassials behind their crushing premolars, therefore they did not need to wait for their prey to die, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's turf. [7] Spotted hyenas spread from their original homeland during the Middle Pleistocene, and quickly colonised a very wide area from Europe, to southern Africa and China. [10] With the decline of grasslands 12,500 years ago, Europe experienced a massive loss of lowland habitats favoured by spotted hyenas, and a corresponding increase in mixed woodlands. Spotted hyenas, under these circumstances, would have been outcompeted by wolves and humans, who were as much at home in forests as in open landsand in highlands as in lowlands. Spotted hyena populations began to shrink after roughly 20,000 years ago, completely disappearing from Western Europe between 11 and 14 thousand years ago, and earlier in some areas. [11]

Genera of the Hyaenidae (extinct and recent)

Reconstruction of Pachycrocuta brevirostris Short-faced hyena1.JPG
Reconstruction of Pachycrocuta brevirostris
A spotted hyena of subfamily Hyaeninae Crocuta crocuta.jpg
A spotted hyena of subfamily Hyaeninae

The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997) [12] and Wozencraft (2005) in Wilson and Reeders Mammal Species of the World for extant genera. [13] The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family Percrocutidae (though they are generally grouped as sister-taxa to hyenas [14] ). Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus Pachycrocuta, but in the genus Hyaena. The Protelinae (aardwolves) are not treated as a separate subfamily, but included in the Hyaeninae.

Phylogeny

The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991), [21] as updated by Turner et al. (2008). [22]

Evolution of hyaenas

Protictitherium crassum

"Protictitherium" cingulatum

"Protictitherium" intermedium

"Protictitherium" llopisi

"Protictitherium" punicum

"Protictitherium" gaillardi

"Protictitherium" sumegense

"Protictitherium" csakvarense

Plioviverrops

Plioviverrops gervaisi

    Civet/mongoose-like insectivore-omnivores

Plioviverrops orbignyi

Plioviverrops guerini

Plioviverrops faventinus

Plioviverrops gaudryi

Tungurictis spocki

Thalassictis robusta

"Thalassictis" certa

"Thalassictis" montadai

"Thalassictis" proava

"Thalassictis" sarmatica

"Thalassictis" spelaea

Tongxinictis primordialis

Proteles

Proteles cristatus (aardwolf) The life of animals (Colored Plate 4) (proteles cristatus).jpg

Proteles amplidentus

Ictitherium

Ictitherium viverrinum

Ictitherium ebu

    Jackal-like hyaenas

Ictitherium tauricum

Ictitherium ibericum

Ictitherium kurteni

Ictitherium intuberculatum

Ictitherium pannonicum

Miohyaenotherium bessarabicum

Hyaenotherium

Hyaenotherium wongii

Hyaenictitherium hyaenoides

"Hyaenictitherium" pilgrimi

"Hyaenictitherium" parvum

"Hyaenictitherium" namaquensis

"Hyaenictitherium" minimum

Lycyaena

Lycyaena chaeretis

Lycyaena dubia

    Cursorial hunting hyaenas

Lycyaena macrostoma

Lycyaena crusafonti

Hyaenictis

Hyaenictis graeca

Hyaenictis almerai

Hyaenictis hendeyi

Lycyaenops

Lycyaenops rhomboideae

Lycyaenops silberbergi

Chasmaporthetes

Chasmaporthetes exitelus

Chasmaporthetes bonisi

Chasmaporthetes borissiaki

Chasmaporthetes lunensis

Chasmaporthetes melei

Chasmaporthetes ossifragus

Chasmaporthetes sp. Florida

Chasmaporthetes nitidula

Chasmaporthetes australis

(running hyaenas)
Hyaeninae
Bone-cracking hyaenas    

Metahyaena confector

Palinhyaena reperta

Hyaenid sp. E Langebaar

Belbus beaumonti

Hyaena abronia

Hyaena hyaena (striped hyaena) Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg

Parahyaena howelli

Parahyaena brunnea (brown hyaena) Hyaena fusca (white background).jpg

Pliocrocuta perrieri

Pachycrocuta brevirostris (giant hyaena) Pachycrocuta brevirostris restoration.jpg

Adcrocuta eximia Adcrocuta eximia restoration.jpg

Allohyaena kadici

Crocuta crocuta (spotted hyaena) Hyaena maculata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg   

Crocuta eturono

(bonecracking hyenas)

Phylogenic relationships based on morphological characteristics, after Werdelin & Solounias (1991) and Turner et al (2008).

A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli et al, 2006 [23] ).

Hyaenidae

Proteles cristatus (aardwolf) The life of animals (Colored Plate 4) (proteles cristatus).jpg

Crocuta crocuta (spotted hyena) Hyaena maculata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg

Parahyaena brunnea (brown hyena) Hyaena fusca (white background).jpg

Hyaena hyaena (striped hyena) Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg

Characteristics

Build

Striped hyena skull. Note the disproportionately large carnassials and premolars adapted for bone consumption StripedHyenaSkull.jpg
Striped hyena skull. Note the disproportionately large carnassials and premolars adapted for bone consumption
Aardwolf skull. Note the greatly reduced molars and carnassials, rendered redundant from insectivory Aardwolfskull.jpg
Aardwolf skull. Note the greatly reduced molars and carnassials, rendered redundant from insectivory

Hyenas have relatively short torsos and are fairly massive and wolf-like in build, but have lower hind quarters, high withers and their backs slope noticeably downward toward their rumps. The forelegs are high, while the hind legs are very short and their necks are thick and short. Their skulls superficially resemble those of large canids, but are much larger and heavier, with shorter facial portions. Hyenas are digitigrade, with the fore and hind paws having four digits each and sporting bulging pawpads. [24] Like canids, hyenas have short, blunt, non-retractable claws. [25] Their pelage is sparse and coarse with poorly developed or absent underfur. Most species have a rich mane of long hair running from the withers or from the head. [24] With the exception of the spotted hyena, hyaenids have striped coats, which they likely inherited from their viverrid ancestors. [7] Their ears are large and have simple basal ridges and no marginal bursa. [25] Their vertebral column, including the cervical region are of limited mobility. Hyenas have no baculum. [26] Hyenas have one more pair of ribs than canids do, and their tongues are rough like those of felids and viverrids. [27] Males in most hyena species are larger than females, [28] though the spotted hyena is exceptional, as it is the female of the species that outweighs and dominates the male. Also, unlike other hyenas, the female spotted hyena's external genitalia closely resembles that of the male. [29]

Their dentition is similar to that of the canid, but is more specialised for consuming coarse food and crushing bones. The carnassials, especially the upper, are very powerful and are shifted far back to the point of exertion of peak pressure on the jaws. The other teeth, save for the underdeveloped upper molars, are powerful, with broad bases and cutting edges. The canines are short, but thick and robust. [26] Labiolingually, their mandibles are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and premolars, unlike canids, which do so with their post-carnassial molars. [30] The strength of their jaws is such that both striped and spotted hyenas have been recorded to kill dogs with a single bite to the neck without breaking the skin. [31] [32] The spotted hyena is renowned for its strong bite proportional to its size, but a number of other animals (including the Tasmanian devil) are proportionately stronger. [33] [34] The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dentition as the other three species. [35] The dental formula for all hyena species is: 3.1.4.13.1.3.1

Although hyenas lack perineal scent glands, they have a large pouch of naked skin located at the anal opening. Large anal glands above the anus open into this pouch. Several sebaceous glands are present between the openings of the anal glands and above them. [25] These glands produce a white, creamy secretion that the hyenas paste onto grass stalks. The odour of this secretion is very strong, smelling of boiling cheap soap or burning, and can be detected by humans several metres downwind. [36] The secretions are primarily used for territorial marking, though both the aardwolf [7] and the striped hyena [37] will spray them when attacked.

Behavior

Spotted hyena cubs at their den 2667 Spotted Hyena Cubs.JPG
Spotted hyena cubs at their den
Brown hyena cub standing on a path of stones Parahyaena brunnea puppy.jpg
Brown hyena cub standing on a path of stones

Hyenas groom themselves often like felids and viverrids, and their way of licking their genitals is very cat-like (sitting on the lower back, legs spread with one leg pointing vertically upward). However, unlike other feliforms, they do not "wash" their faces. They defecate in the same manner as other Carnivora, though they never raise their legs as canids do when urinating, as urination serves no territorial function for them. Instead, hyenas mark their territories using their anal glands, a trait found also in viverrids and mustelids, but not canids and felids. [38] When attacked by lions or dogs, striped [39] and brown hyenas [40] will feign death, though the spotted hyena will defend itself ferociously. [32] The spotted hyena is very vocal, producing a number of different sounds consisting of whoops, grunts, groans, lows, giggles, yells, growls, laughs and whines. [41] The striped hyena is comparatively silent, its vocalisations being limited to a chattering laugh and howling. [42]

Whoop of a spotted hyena in Umfolosi Game Park, South Africa.

Mating between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally engage in a single, drawn out copulation. [38] Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings. [43] In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears. [44] Hyenas do not regurgitate food for their young and male spotted hyenas play no part in raising their cubs, [38] though male striped hyenas do so. [45]

The striped hyena is primarily a scavenger, though it will also attack and kill any animals it can overcome, [39] and will supplement its diet with fruit. [46] The spotted hyena, though it also scavenges occasionally, is an active pack hunter of medium to large sized ungulates, which it catches by wearing them down in long chases and dismembering them in a canid-like manner. The aardwolf is primarily an insectivore, specialised for feeding on termites of the genus Trinervitermes and Hodotermes , which it consumes by licking them up with its long, broad tongue. An aardwolf can eat 300,000 Trinervitermes on a single outing. [7] Spotted hyenas may kill as many as 95% of the animals they eat, [47] while striped hyenas are largely scavengers. [39] Generally, hyenas are known to drive off larger predators, like lions, from their kills, despite having a reputation in popular culture for being cowardly. [39] Hyenas are primarily nocturnal animals, but sometimes venture from their lairs in the early-morning hours. With the exception of the highly social spotted hyena, hyenas are generally not gregarious animals, though they may live in family groups and congregate at kills. [48] Spotted hyenas are one of the few mammals other than bats known to survive infection with rabies virus. [49] and have showed little or no disease-induced mortality during outbreaks in sympatric carnivores. Despite this perceived unique disease resistance, little is known about the immune system of spotted hyenas, [50] [51] [52] [53] and even less is know about other Hyaenidae species.

Relationships with humans

Folklore, mythology and literature

Cave hyena (Crocuta crocuta spelaea) painting found in the Chauvet Cave in 1994 20,000 Year Old Cave Paintings Hyena.png
Cave hyena (Crocuta crocuta spelaea) painting found in the Chauvet Cave in 1994
A striped hyena, as depicted on the Nile mosaic of Palestrina Hyenamosaic.jpg
A striped hyena, as depicted on the Nile mosaic of Palestrina

Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names. In western African tales, spotted hyenas are sometimes depicted as bad Muslims who challenge the local animism that exists among the Beng in Côte d’Ivoire.[ citation needed ] In East Africa, Tabwa mythology portrays the spotted hyena as a solar animal that first brought the sun to warm the cold earth, while West African folklore generally shows the hyena as symbolising immorality, dirty habits, the reversal of normal activities, and other negative traits. In Tanzania, there is a belief that witches use spotted hyenas as mounts. [54] In the Mtwara Region of Tanzania, it is believed that a child born at night while a hyena is crying will likely grow up to be a thief. In the same area, hyena faeces are believed to enable a child to walk at an early age, thus it is not uncommon in that area to see children with hyena dung wrapped in their clothes. [55] The Kaguru of Tanzania and the Kujamaat of Southern Senegal view hyenas as inedible and greedy hermaphrodites. A mythical African tribe called the Bouda is reputed to house members able to transform into hyenas. [56] A similar myth occurs in Mansôa. These "werehyenas" are executed when discovered, but do not revert to their human form when killed. [55]

Striped hyenas are often referred to in Middle Eastern literature and folklore, typically as symbols of treachery and stupidity. [57] In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of jinns. [54] Arab writer al-Qazwīnī (1204–1283) spoke of a tribe of people called al-Ḍabyūn meaning "hyena people". In his book ‘Ajā’ib Al-Makhlūqāt he wrote that should one of this tribe be in a group of 1000 people, a hyena could pick him out and eat him. [57] A Persian medical treatise written in 1376 tells how to cure cannibalistic people known as kaftar, who are said to be “half-man, half-hyena”. [54] al-Damīrī in his writings in Ḥawayān al-Kubrā (1406) wrote that striped hyenas were vampiric creatures that attacked people at night and sucked the blood from their necks. He also wrote that hyenas only attacked brave people. Arab folklore tells of how hyenas can mesmerise victims with their eyes or sometimes with their pheromones. [57] In a similar vein to al-Damīrī, the Greeks, until the end of the 19th century, believed that the bodies of werewolves, if not destroyed, would haunt battlefields as vampiric hyenas that drank the blood of dying soldiers. [58] The image of striped hyenas in Afghanistan, India and Palestine is more varied. Though feared, striped hyenas were also symbolic of love and fertility, leading to numerous varieties of love medicine derived from hyena body parts. Among the Baluch and in northern India, witches or magicians are said to ride striped hyenas at night. [54]

The striped hyena is mentioned in the Bible. The Arab word for the hyena, ḍab` or ḍabu` (plural ḍibā`), is alluded to in a valley in Israel known as Shaqq-ud-Diba` (meaning "cleft of the hyenas") and Wadi-Abu-Diba` (meaning "valley of the hyenas"). Both places have been interpreted by some scholars as being the Biblical Valley of Tsebo`im mentioned in 1 Samuel 13:18. The modern Hebrew word for hyena is tzavoa`, which is similar to the word "tsavua`" meaning "colored". Though the Authorized King James Version of the Bible interprets the term "`ayit tsavua`" (which appears in Jeremiah 12:9) as "speckled bird", Henry Baker Tristram argued that it was most likely a hyena being mentioned. [59]

The vocalisation of the spotted hyena resembling hysterical human laughter has been alluded to in numerous works of literature: "to laugh like a hyæna" was a common simile, and is featured in The Cobbler's Prophecy (1594), Webster's Duchess of Malfy (1623) and Shakespeares As You Like It , Act IV. Sc.1.

Die Strandjutwolf (The brown hyena) is an allegorical poem by the renowned South African poet, N. P. van Wyk Louw, which evokes a sinister and ominous presence.

Bud and Lou, from the DC Comics, are also hyenas and are pets of Harley Quinn.

Attacks on humans

Illustration from Fraser's magazine showing an artist's impression of a "stag-hound" biting a spotted hyena attacking its master Hyena attack (1842 woodcut).jpg
Illustration from Fraser's magazine showing an artist's impression of a "stag-hound" biting a spotted hyena attacking its master
A 1739 advertisement by Charles Benjamin Incledon featuring feliforms: the Mesopotamian lion from the vicinity of Bassorah, Cape lion, tiger from the East Indies, panther from Buenos Aires, Hyaena hyaena from West Africa, and leopard from Turkey, besides a "Man tyger" from Africa. The advertisement mentions that the 'hyaena' can mimic a human voice to lure humans. Bodleian Libraries, Handbill of Merchant's Hall, 1739, announcing A lion, lionesses, tigers, etc..jpg
A 1739 advertisement by Charles Benjamin Incledon featuring feliforms: the Mesopotamian lion from the vicinity of Bassorah, Cape lion, tiger from the East Indies, panther from Buenos Aires, Hyaena hyaena from West Africa, and leopard from Turkey, besides a "Man tyger" from Africa. The advertisement mentions that the 'hyaena' can mimic a human voice to lure humans.

Among hyenas, only the spotted and striped hyenas have been known to become man-eaters. Hyenas are known to have preyed on humans in prehistory: Human hair has been found in fossilised hyena dung dating back 195,000 to 257,000 years. [60] Some paleontologists believe that competition and predation by cave hyenas (Crocuta crocuta spelaea) in Siberia was a significant factor in delaying human colonisation of Alaska. Hyenas may have occasionally stolen human kills, or entered campsites to drag off the young and weak, much like modern spotted hyenas in Africa. The oldest Alaskan human remains coincide with roughly the same time cave hyenas became extinct, leading certain paleontologists to infer that hyena predation was what prevented humans crossing the Bering strait earlier. [61] Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup [62] and the Red Terror. [63] Hyenas habituated to scavenging on human corpses may develop bold behaviours towards living people; hyena attacks on people in southern Sudan increased during the Second Sudanese Civil War, when human corpses were readily available to them. [64]

Although spotted hyenas do prey on humans in modern times, such incidents are rare. However, according to the SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique), attacks on humans by spotted hyenas are likely to be underreported. [65] According to hyena expert Dr. Hans Kruuk, man-eating spotted hyenas tend to be very large specimens: A pair of man-eating hyenas, responsible for killing 27 people in Mulanje (Mlanje), Malawi in 1962, were weighed at 72 kg (159 lb) and 77 kg (170 lb) after being shot. [66] In 1903, Hector Duff wrote of how spotted hyenas in the Mzimba district of Angoniland would wait at dawn outside people's huts and attack them when they opened their doors. [67] Victims of spotted hyenas tend to be women, children and sick or infirm men: Theodore Roosevelt wrote on how in 1908–1909 in Uganda, spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps. [68] Spotted hyenas are widely feared in Malawi, where they have been known to occasionally attack people at night, particularly during the hot season when people sleep outside. Hyena attacks were widely reported in Malawi's Phalombe plain, to the north of Michesi Mountain. Five deaths were recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961 when eight people were killed. Attacks occurred most commonly in September, when people slept outdoors, and bush fires made the hunting of wild game difficult for the hyenas. [65] [67] An anecdotal news report from the World Wide Fund for Nature 2004 indicates that 35 people were killed by spotted hyenas in a 12-month period in Mozambique along a 20 km stretch of road near the Tanzanian border. [65]

In ordinary circumstances, striped hyenas are extremely timid around humans, though they may show bold behaviours toward people at night. [69] On rare occasions, striped hyenas have preyed on humans. In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the Iğdır Province, with 25 children and 3 adults being wounded in one year. The attacks provoked local authorities into announcing a reward of 100 rubles for every hyena killed. Further attacks were reported later in some parts of Transcaucasia, particularly in 1908. Instances are known in Azerbaijan of striped hyenas killing children sleeping in courtyards during the 1930s and 1940s. In 1942, a sleeping guard was mauled in his hut by a hyena in Qalıncaq (Golyndzhakh). Cases of children being taken by hyenas by night are known in southeast Turkmenia's Bathyz Nature Reserve. A further attack on a child was reported around Serakhs in 1948. [70] Several attacks have occurred in India; in 1962, nine children were thought to have been taken by hyenas in the town of Bhagalpur in the Bihar State in a six-week period [59] and 19 children up to the age of four were killed by hyenas in Karnataka in 1974. [71] A consensus on wild animal attacks during a five-year period in the Indian state of Madhya Pradesh showed that hyenas had only attacked three people, the lowest figure when compared to deaths caused by wolves, gaur, boar, elephants, tigers, leopards and sloth bears. [72]

Hyenas as food and medicine

Hyenas are used for food and medicinal purposes in Somalia [73] although it is considered haraam in Islam. [74] This practice dates back to the times of the Ancient Greeks and Romans, who believed that different parts of the hyena's body were effective means to ward off evil and to ensure love and fertility. [54]

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Cenozoic Third era of the Phanerozoic Eon 66 million years ago to present

The Cenozoic Era meaning "new life" is the current and most recent of the three geological eras of the Phanerozoic Eon. It follows the Mesozoic Era and extends from 66 million years ago to the present day. It is generally believed to have started on the first day of the Cretaceous–Paleogene extinction event when an asteroid hit the Earth.

Canidae Family of mammals

Canidae is a biological family of dog-like carnivorans. A member of this family is called a canid. There are three subfamilies found within the canid family, which are the extinct Borophaginae and Hesperocyoninae, and the extant Caninae. The Caninae are known as canines, which includes domestic dogs, wolves, foxes, and other extant and extinct species.

<i>Hyaena</i> genus of mammals in the order Carnivora

Hyaena is a genus comprising two of the living species of hyenas: the striped hyena from Western Asia, India, Central Asia, East Africa and Northern Africa and the brown hyena from Southern Africa. The brown hyena has sometimes been placed in a separate genus Parahyaena, or even included in the otherwise fossil genus Pachycrocuta, but recent sources have tended to place it in Hyaena.

<i>Vulpes</i>

Vulpes is a genus of the dog subfamily Caninae. The members of this genus are colloquially referred to as true foxes, meaning they form a proper clade. The word "fox" occurs on the common names of species. True foxes are distinguished from members of the genus Canis, such as domesticated dogs, wolves, jackals and coyotes, by their smaller size (5–11 kg) and flatter skull. They have black, triangular markings between their eyes and nose, and the tip of their tail is often a different color from the rest of their pelt. The typical lifespan for this genus is between two and four years, but can reach up to a decade.

<i>Canis</i>

Canis is a genus of the Caninae containing multiple extant species, such as wolves, dogs, coyotes and jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.

Striped hyena Species of hyena

The striped hyena is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia and the Indian subcontinent. It is listed by the IUCN as near-threatened, as the global population is estimated to be under 10,000 mature individuals which continues to experience deliberate and incidental persecution along with a decrease in its prey base such that it may come close to meeting a continuing decline of 10% over the next three generations. It is also the national animal of Lebanon.

Brown hyena

The brown hyena, also called strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is currently the rarest species of hyena. The largest remaining brown hyena population is located in the southern Kalahari Desert and coastal areas in Southwest Africa. The global population of brown hyena is estimated by IUCN at a number between 4,000 and 10,000 and its conservation status is marked as near threatened in the IUCN Red List.

Spotted hyena Species of hyena

The spotted hyena, also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus Crocuta, native to sub-Saharan Africa. It is listed as being of least concern by the IUCN on account of its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. The species may have originated in Asia, and once ranged throughout Europe for at least one million years until the end of the Late Pleistocene. The spotted hyena is the largest known member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear-like build, its rounded ears, its less prominent mane, its spotted pelt, its more dual purposed dentition, its fewer nipples and the presence of a pseudo-penis in the female. It is the only mammalian species to lack an external vaginal opening.

Caniformia

Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include dogs, bears, wolves, foxes, raccoons, badgers, seals and mustelids. The Pinnipedia are also assigned to this group. The center of diversification for Caniformia is North America and northern Eurasia. Caniformia stands in contrast to the other suborder of Carnivora, Feliformia, the center of diversification of which was in Africa and southern Asia.

<i>Pachycrocuta</i> Genus of mammals (fossil)

Pachycrocuta was a genus of prehistoric hyenas. The largest and most well-researched species is Pachycrocuta brevirostris, colloquially known as the giant short-faced hyena as it stood about 90–100 cm (35–39 in) at the shoulder and it is estimated to have averaged 110 kg (240 lb) in weight, approaching the size of a lioness, making it the largest known hyena. Pachycrocuta first appeared during the late Miocene and became extinct during the middle Pleistocene, 400,000 years ago.

<i>Chasmaporthetes</i>

Chasmaporthetes, also known as hunting or running hyena, is an extinct genus of hyenas distributed in Eurasia, North America, and Africa during the Pliocene-Pleistocene epochs, living from 4.9 million to 780,000 years ago, existing for about 4.12 million years . The genus probably arose from Eurasian Miocene hyenas such as Thalassictis or Lycyaena, with C. borissiaki being the oldest known representative. The species C. ossifragus was the only hyena to cross the Bering land bridge into the Americas, and ranged over what is now Arizona and Mexico during Blancan and early Irvingtonian Land Mammal ages, between 5.0 and 1.5 million years ago.

Cave hyena

The cave hyena, also known as the Ice Age spotted hyena, was a paleosubspecies of spotted hyena which ranged from the Iberian Peninsula to eastern Siberia. It is one of the best known mammals of the Ice Age and is well represented in many European bone caves. The cave hyena was a highly specialised animal, with its progressive and regressive features being more developed than in its modern African relative. It preyed on large mammals, and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal caves, sinkholes, mud pits and muddy areas along rivers.

Feliformia

Feliformia is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats, hyenas, mongooses, viverrids, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Caniformia.

Caninae

The Caninae, known as canines, are one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Borophaginae and Hesperocyoninae. The Caninae includes all living canids and their most recent fossil relatives. Their fossils were first found in North America and dated to the Oligocene era, then spreading to Asia at the end of the Miocene era, some 7 million to 8 million years ago.

<i>Xenocyon</i> sub genus of extinct carnivores

Xenocyon is an extinct subgenus of Canis. The group includes Canis (Xenocyon) africanus, Canis (Xenocyon) antonii and Canis (Xenocyon) falconeri that gave rise to Canis (Xenocyon) lycanoides. The hypercarnivore Xenocyon gave rise to the modern dhole and the African wild dog.

<i>Amphimachairodus</i> genus of mammals

Amphimachairodus is an extinct genus of large machairodonts belonging to the clade known as Eumachairodontia along with relatives like Smilodon and Homotherium. It is also a member of the tribe Homotherini within Machairodontidae and is most closely related to such species as Xenosmilus, Homotherium itself, and Nimravides. It inhabited Eurasia, Northern Africa and North America during the late Miocene epoch.

African golden wolf

The African golden wolf is a canine native to North Africa and the Horn of Africa. It is the descendant of a genetically admixed canid of 72% gray wolf and 28% Ethiopian wolf ancestry. It occurs in Senegal, Nigeria, Chad, Morocco, Algeria, Tunisia, Libya, Kenya, Egypt, and Tanzania. It is listed as least concern on the IUCN Red List. In the Atlas Mountains, it was sighted in elevations as high as 1,800 m (5,900 ft). It is primarily a predator, targeting invertebrates and mammals as large as gazelle fawns, though larger animals are sometimes taken. Its diet also includes animal carcasses, human refuse, and fruit. The African golden wolf is a monogamous and territorial species; offspring remain with the family to assist in raising their parents' younger pups.

Ussuri dhole subspecies of mammal

The Ussuri dhole, also known as the Indian dhole, Eastern Asiatic dhole, Chinese dhole or Southern dhole, is the nominate subspecies of the dhole native to East Asia. It is widespread in the Indian subcontinent and the Indochinese Peninsula. The Ussuri dhole is also native to China, however it is probably extinct in most of its ranges in China, as well as in Mongolia and the Russian Far East.

Herpestoidea Superfamily of mammals

Herpestoidea is a superfamily of mammalia carnivores which includes mongooses, Malagasy carnivorans and the hyenas.

References

Notes

  1. ὕαινα, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus. Etymologically, this is a feminine of ὕς "swine".
  2. Wilson, D.E.; Mittermeier, R.A., eds. (2009). Handbook of the Mammals of the World, Volume 1: Carnivora. Barcelona: Lynx Edicions. pp. 50–658. ISBN   978-84-96553-49-1.
  3. Mills & Hofer 1998 , p. iv
  4. Kruuk 1972 , p. 274
  5. Mills & Hofer 1998 , p. 96
  6. Vats, Rajeev; Thomas, Simion (7 May 2015). "A study on use of animals as traditional medicine by Sukuma Tribe of Busega District in North-western Tanzania". Journal of Ethnobiology and Ethnomedicine. 11: 38. doi:10.1186/s13002-015-0001-y. PMC   4472419 . PMID   25947365.
  7. 1 2 3 4 5 6 7 8 9 Macdonald 1992 , pp. 119–144
  8. Kurtén 1968 , pp. 66–68
  9. Mills & Hofer 1998 , p. 1
  10. 1 2 Kurtén 1968 , pp. 69–72
  11. "Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy" (PDF). C. Stiner, Mary. Revue de Paléobiologie, Genève. Retrieved 2008-09-16.
  12. Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level, Columbia University Press, New York 1997, 631 Seiten, ISBN   0-231-11013-8
  13. Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–548. ISBN   978-0-8018-8221-0. OCLC   62265494.
  14. Figueirido, Borja; Jack Tseng, Zhijie; Martín-Serra, Alberto (2013). "Skull Shape Evolution In Durophagous Carnivorans". Evolution. 67 (7): 1975–93. doi:10.1111/evo.12059. PMID   23815654. S2CID   23918004.
  15. Coca-Ortega, C.; Pérez-Claros, J.A. (January 2019). "Characterizing ecomorphological patterns in hyenids: a multivariate approach using postcanine dentition". PeerJ. 6: 6:e6238. doi: 10.7717/peerj.6238 . PMC   6330948 . PMID   30648005.
  16. Werdelin, Lars (March 1996). "Community‐wide character displacement in Miocene hyaenas". Lethaia. 29 (1): 97–106. doi:10.1111/j.1502-3931.1996.tb01843.x.
  17. Wang, X.; Tsang, Z.J.; et al. (February 2020). "A new species of Tungurictis Colbert, 1939 (Carnivora, Hyaenidae) from the middle Miocene of Junggar Basin, northwestern China and the early divergence of basal hyaenids in East Asia". Geodiversitas. 42 (3): 29–45. doi: 10.5252/geodiversitas2020v42a3 .
  18. 1 2 Turner, A.; Antón, M.; Werdelin, L. (September 2008). "Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe". Geobios. 41 (5): 677–687. doi:10.1016/j.geobios.2008.01.001.
  19. Tseng, Z.J.; Stynder, D. (March 2011). "Mosaic functionality in a transitional ecomorphology: skull biomechanics in stem Hyaeninae compared to modern South African carnivorans". Biological Journal of the Linnean Society. 102 (3): 540–559. doi: 10.1111/j.1095-8312.2010.01602.x .
  20. Vinuesa, V.; Madurell-Malapeira, J.; et al. (April 2016). "A new skull of Hyaenictis Gaudry, 1861 (Carnivora, Hyaenidae) shows incipient adaptations to durophagy". Journal of Mammalian Evolution. 24 (2): 207–219. doi:10.1007/s10914-016-9334-0. S2CID   23453043.
  21. Werdelin, L.; Solounias, 1991 (1991). "The Hyaenidae: taxonomy, systematics and evolution" (PDF). Fossils and Strata. 30: 1–104.CS1 maint: numeric names: authors list (link)
  22. Turner, Alan; Antón, Mauricio; Werdelin, Lars (2008). "Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe". Geobios. 41 (5): 677–687. doi:10.1016/j.geobios.2008.01.001.
  23. Koepfli, K.-P.; Jenks, S. M.; Eizirik, E.; Zahirpour, T.; Van Valkenburgh, B.; Wayne, R. K. (2006). "Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix". Molecular Phylogenetics and Evolution. 38 (3): 603–620. doi:10.1016/j.ympev.2005.10.017. PMID   16503281.
  24. 1 2 Heptner & Sludskii 1992 , p. 3
  25. 1 2 3 Pocock 1941 , pp. 62–63
  26. 1 2 Heptner & Sludskii 1992 , pp. 4–5
  27. Holl, William & Wood, Neville The Analyst: a quarterly journal of science, literature, natural history, and the fine arts, Volume 10, p. 59, Simpkin & Marshall, 1840
  28. Mills & Hofer 1998 , p. 21
  29. Kruuk 1972 , pp. 210–211
  30. Therrien, François (2005). "Mandibular force profiles of extant carnivorans and implications for the feeding behavior of extinct predators". Journal of Zoology. 267 (3): 249–270. doi:10.1017/S0952836905007430.
  31. Daniel Johnson (1827) Sketches of Indian Field Sports: With Observations on the Animals; Also an Account of Some of the Customs of the Inhabitants; with a Description of the Art of Catching Serpents, as Practiced by the Conjoors and Their Method of Curing Themselves when Bitten: with Remarks on Hydrophobia and Rabid Animals p. 45-46, R. Jennings, 1827
  32. 1 2 Stevenson-Hamilton, James (1917) Animal life in Africa, Vol. 1, p.95, London : William Heinemann
  33. Salleh, Anna (4 April 2005). "Marsupial has the deadliest bite". abc.net.au. Retrieved 24 January 2013.
  34. Wroe, S.; McHenry, C.; Thomason, J. (2005). "Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa". Proceedings of the Royal Society B: Biological Sciences. 272 (1563): 619–625. doi:10.1098/rspb.2004.2986. PMC   1564077 . PMID   15817436.
  35. Richardson, Philip K.R.; Bearder, Simon (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp.  154–159. ISBN   0-87196-871-1.
  36. Kruuk 1972 , p. 222
  37. Heptner & Sludskii 1992 , p. 38
  38. 1 2 3 Kruuk 1972 , pp. 271–73
  39. 1 2 3 4 Pocock 1941 , p. 72
  40. Mills & Mills 2010 , pp. 60–61
  41. Kruuk 1972 , p. 220
  42. Pocock 1941 , p. 73
  43. Kruuk 1972 , pp. 247–249
  44. Rosevear 1974 , p. 350
  45. Heptner & Sludskii 1992 , pp. 40–42
  46. Heptner & Sludskii 1992 , pp. 31–33
  47. Hyaena Specialist Group - Spotted Hyena: Diet and Foraging Archived 2011-02-04 at the Wayback Machine . Hyaenidae.org. Retrieved on 2015-11-06.
  48. Rosevear 1974 , pp. 343–344
  49. East, M.L. (18 December 2001). "Regular exposure to rabies virus and lack of symptomatic disease in Serengeti spotted hyenas". PNAS. 98 (26): 15026–31. doi:10.1073/pnas.261411898. PMC   64977 . PMID   11742089.
  50. Flies, A.S.; et al. (29 February 2016). "Socioecological predictors of immune defenses in a wild spotted hyenas". Functional Ecology. 30 (9): 1549–1557. doi:10.1111/1365-2435.12638. PMC   5098940 . PMID   27833242.
  51. Flies, A.S.; et al. (7 October 2015). "Markedly Elevated Antibody Responses in Wild versus Captive Spotted Hyenas Show that Environmental and Ecological Factors Are Important Modulators of Immunity". PLOS ONE. 10 (10): e0137679. doi:10.1371/journal.pone.0137679. PMC   4621877 . PMID   26444876.
  52. Flies, A.S.; et al. (15 January 2012). "Development of a hyena immunology toolbox". Veterinary Immunology and Immunopathology. 145 (1–2): 110–9. doi:10.1016/j.vetimm.2011.10.016. PMC   3273618 . PMID   22173276.
  53. Flies, A.S.; et al. (2 February 2014). "Characterization of toll-like receptors 1-10 in spotted hyenas". Veterinary Research Communications. 38 (2): 165–70. doi:10.1007/s11259-014-9592-3. PMC   4112752 . PMID   24488231.
  54. 1 2 3 4 5 Frembgen, Jürgen W. The Magicality of the Hyena: Beliefs and Practices in West and South Asia , Asian Folklore Studies, Volume 57, 1998: 331–344
  55. 1 2 Mills & Hofer 1998 , p. 97
  56. "The spotted hyena from Aristotle to the Lion King: reputation is everything - In the Company of Animals". Stephen E. Glickman. Archived from the original on 2011-08-11. Retrieved 2007-05-22.
  57. 1 2 3 Mounir R. Abi-Said (2006) Reviled as a grave robber: The ecology and conservation of striped hyaenas in the human dominated landscapes of Lebanon Ph.D. thesis, University of Kent (Biodiversity management)
  58. Woodward, Ian (1979). The Werewolf Delusion. p. 256. ISBN   0-448-23170-0.
  59. 1 2 Bright, Michael (2006). Beasts of the Field: The Revealing Natural History of Animals in the Bible. pp.  127–129. ISBN   1-86105-831-4.
  60. ''Oldest Human Hair Found in Fossilized Dung'' Jennifer Viegas, Discovery News. Dsc.discovery.com (2009-02-10). Retrieved on 2015-11-06.
  61. "Hyenas and Humans in Ice Age Siberia" (PDF). Christy G. Turner II. School of Human Evolution and Social Change, Arizona State University. Retrieved 2008-08-02.[ dead link ]
  62. Kapuściński, Ryszard, The Emperor: Downfall of an Autocrat . 1978. ISBN   0-679-72203-3
  63. Donham, Donald Lewis (1999) Marxist modern: an ethnographic history of the Ethiopian revolution, University of California Press, page 135, ISBN   0-520-21329-7
  64. Copson, Raymond W. (1994) Africa's wars and prospects for peace, M.E. Sharpe, page 6, ISBN   1-56324-300-8
  65. 1 2 3 Begg, Colleen, Begg, Keith & Muemedi, Oscar (2007) Preliminary data on human - carnivore conflict in Niassa National Reserve, Mozambique, particularly fatalities due to lion, spotted hyaena and crocodile Archived 2011-12-26 at the Wayback Machine , SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique)
  66. Kruuk, Hans (2002) Hunter and hunted: relationships between carnivores and people Cambridge University Press, ISBN   0-521-89109-4
  67. 1 2 Knight, John (2000). Natural Enemies: People-Wildlife conflicts in Anthropological Perspective. ISBN   0-415-22441-1.
  68. Roosevelt, Theodore (1910) African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist, New York, C. Scribner's sons
  69. Heptner & Sludskii 1992 , p. 36
  70. Heptner & Sludskii 1992 , p. 46
  71. Mills & Hofer 1998 , p. 25
  72. "The Fear of Wolves: A Review of Wolf Attacks on Humans" (PDF). Norsk Institutt for Naturforskning. Archived from the original (PDF) on 2007-09-27. Retrieved 2008-06-26.
  73. "Islamists authorise hyena meat in Southern Somalia". Somalilandpress. August 12, 2012. Archived from the original on 2017-01-03. Retrieved 2017-01-03.
  74. Agic, Shaykh Senad. "Halal Animals". Eshaykh.com. Archived from the original on 3 January 2017. Retrieved 3 January 2017.

Bibliography

Further reading